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QP91  .H7S^'''°°°'uLin.esuga,   SUGAR  IN  THE  BLOOD  OF  PIGEONS 


RECAP 


BY 

HANNAH  ELIZABETH  HONEYWELL 


A  DISSERTATION 

Submitted  in  Partial  Fulfillment  of  the  Requirements  for  the  Degree 

OF  Doctor  of  Philosophy  in  the  Faculty  of  Pure  Science, 

Columbia  University,  New  York  City 


1921 


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CoIIese  of  Ij^^psimns  mis  burgeons 
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STUDIES  IN  THE  SUGAR  IN  THE  BLOOD  OF  PIGEONS, 


BY 

HANNAH  ELIZABETH  HONEYWELL 


A  DISSERTATION 

Submitted  in  Partial  Fulfillment  of  the  Requirements  for  the  Degree 

OF  Doctor  of  Philosophy  in  the  Faculty  of  Pure  Science, 

Columbia  University,  New  York  City 


1921 


Reprinted  from  The  American  Journal  of  Physiology 
Vol.  58,  No.  1,  November,  1921 


W7r 


STUDIES  OF  THE  SUGAR  IN  THE  BLOOD  OF  PIGEONS 

HANNAH  ELIZABETH  HONEYWELL 
From  the  Department  of  Physiology,  Columbia  University 

Received  for  publication  July  1,  1921 

WTiile  it  is  a  recognized  fact  that  much  of  our  experimental  data 
in  physiology  must  be  obtained  from  animals  other  than  man,  there 
has  been  very  little  hesitation  on  the  part  of  many  experimenters  in 
drawing  conclusions  concerning  phenomena  in  man  from  data  derived 
from  other  animals.  In  many  cases  the  results  have  warranted  the 
practice,  in  others  disappointment  has  resulted.  Actually  the  larger 
the  number  of  experiments  performed,  and  the  greater  the  number  of 
species  from  which  the  data  have  been  derived,  the  more  justification 
there  is  for  a  generalized  statement,  and  for  its  application  to  species 
other  than  those  upon  which  work  has  been  done.  With  this  idea 
in  mind  it  was  thought  wise  to  make  the  determinations  leading  to  the 
data  published  in  the  present  paper,  on  an  animal  hitherto  little  used 
in  metaboUsm  work.  A  bird  rather  than  a  mammal  was  chosen 
largely  because  of  the  fact  that  in  this  form  the  red  blood  corpuscles  are 
nucleated.  That  this  makes  a  striking  difference  in  the  physiology  of 
the  blood  is  well  illustrated  by  Warburg's  (1)  work  on  oxygen  consump- 
tion in  the  drawn  blood  of  birds  as  compared  with  that  of  mammals. 
The  fact  that  the  red  corpuscles  of  birds  are  nucleated  should  prove  of 
especial  value  in  studies  made  on  them  with  the  idea  of  using  the  re- 
sults as  a  means  of  giving  a  better  understanding  of  the  physiology  of 
the  cells  of  tissues  other  than  blood.  A  considerable  literature  has 
developed  upon  the  exchange  of  material  between  the  red  cells  and  the 
plasma.  This  is  especially  true  of  the  chlorides  and  the  sugars.  The 
data  upon  which  this  literature  is  based  have  been  obtained  from 
studies  on  the  blood  of  mammals.  It  seems  probable  that  parallel 
studies  on  bird 's  blood  with  its  nucleated  cells  should  lead  to  a  better 
understanding  of  this  process. 

The  fact  that  the  birds,  characteristically,  have  a  higher  temperature 
than  mammals  is  an  additional  reason  why  their  metabolism  should  be 
studied  for  comparison  with  that  of  mammals.     In  fact,  it  has  been 

152 


BLOOD  SUGAR  OF  PIGEONS  153 

suggested  by  Bierry  (2)  that  this  is  the  reason  for  the  very  high  con- 
centration of  sugar  found  in  their  blood  as  compared  with  that  found 
in  the  blood  of  mammals.     That  there  is  a  correlation  between  body' 
temperature  and  blood  sugar  concentration  is  shown  by  the  work  of 
Hollinger  (3)  and  others. 

Since  most  of  these  properties  are  of  general  interest  the  bird  should 
be  an  instructive  addition  to  our  list  of  laboratory  animals.  Com- 
paratively little  has  been  done  on  the  metabolism  of  birds  aside  from 
some  work  on  polyneuritis.  It  is  therefore  advisable  to  extend  our 
knowledge  of  this  phase  of  their  physiology. 

From  birds  in  general  the  pigeon  was  selected  and  it  is  thought 
that  it  should  be  a  practical  and  convenient  bird  with  which  to  work 
for  the  following  reasons:  a,  it  is  of  convenient  size;  h,  it  is  comparatively 
easy  to  obtain;  c,  its  first  cost  is  not  prohibitive;  d,  it  is  easily  and 
economically  kept;  e,  it  is  a  seed  eater  and  therefore  herbivorous. 
Further,  in  the  use  of  the  carrier  pigeon  there  is  an  opportunity  for  the 
study  of  fatigue.  Mosso  (4),  in  fact,  has  already  made  some  use  of  it 
for  this  purpose.  This  phase  of  the  work  we  hope  to  extend,  and  con- 
sequently further  data  on  resting  ])irds  are  necessary  for  comparison 
with  those  obtained  from  fatigued  birds  and  from  resting  and  fatigued 
mammals. 

Method:  1.  The  care  of  the  birds.  The  pigeons  were  confined  in 
the  laboratory  until  they  became  accustomed  to  their  surroundings. 
They  were  subjected  to  frequent  handling  in  order  to  reduce  their  fear 
of  the  operator,  and  so  to  minimize  as  far  as  possible  results  which  might 
arise  from  fright.  In  most  cases  the  birds  wholly  ceased  to  resist 
handling  when  taken  from  the  cage.  During  periods  between  experi- 
ments the  pigeons  were  kept  in  a  large  enclosure  giving  them  oppor- 
tunity to  fly  about  freely.  The  food  consisted  of  a  mixture  of  com, . 
oats,  barley  and  some  other  grains,  and  a  plentiful  supply  of  fresh 
water.  Access  was  given  to  both  food  and  water  at  all  times  not 
otherwise  noted. 

2.  Method  of  estimation  of  sugar.  MacLean's  (5)  micro-method  was 
used  throughout  the  research.  The  macro-method  had  been  success- 
fully used  in  this  laboratory  for  other  blood  sugar  determinations. 
The  micro-method  possesses  the  obvious  advantage  of  requiring  only  a 
small  quantity  of  blood  (0.2  cc).  It  therefore  may  be  used  for  animals 
having  a  comparatively  small  amount  of  blood,  and  it  permits  of 
drawing  consecutive  samples  at  relatively  short  intervals,  without  pro- 
ducing serious  effects  from  hemorrhage.     The  extreme  limits  of  error 


154  HANNAH    ELIZABETH    HONEYWELL 

were  found  to  be  about  10  mgm.  per  100  cc.  of  blood.  Variations  of 
this  amount  or  less  may,  therefore,  be  disregarded. 

3.  Special  teGhniqiie.  While  the  samples  of  blood  were  being  drawn 
the  birds  were  encased  in  a  strong  cloth  jacket  made  especially  for  the 
purpose.  This  was  done  as  a  matter  of  convenience  to  the  operator 
and  of  safety  to  the  birds.  The  jacket  laced  across  the  ventral  surface 
of  the  body  in  such  a  manner  as  to  be  adjustable  to  birds  of  varying 
size,  and  to  secure  the  wings  and  legs. 

The  blood  was  drawn  directly  from  the  heart  into  a  0.2  cc.  pipette 
by  means  of  a  long  hypodermic  needle.  This  needle  was  inserted  at 
the  end  of  the  breast  bone,  going  directly  through  the  skin  and  body  wall 
into  the  abdominal  cavity.  The  tip  of  the  needle  followed  the  breast 
bone  up  to  the  region  of  the  heart,  and  so  avoided  puncture  of  the  liver 
and  other  viscera.  When  the  heart  could  be  felt  at  the  tip  of  the 
needle  the  point  was  dropped  slightly  and  plunged  into  the  heart  tissue. 
Following  this  procedure  the  needle  struck  the  heart  in  such  a  way 
that  arterial  blood  was  drawn.  Post-mortem  examination  showed 
that  the  needle  usually  entered  the  heart  near  the  apex  on  the  left 
side.  Hj^podermic  needles,  20  gauge,  and  3  inches  long,  were  used 
most  successfully.  Since  the  distance  from  the  tip  of  the  breast  bone 
to  the  heart  is  very  neaily  equal  to  the  length  of  such  a  needle,  it  is 
obvious  that  this  method  of  drawing  blood  would  not  be  practicable 
for  birds  much  larger  than  the  pigeon.  It  is  also  probable,  because  of 
the  lesser  quantity  of  blood  and  the  smaller  and  more  delicate  heart, 
that  this  method  would  not  be  applicable  to  species  much  smaller. 
All  the  experiments  were  performed  during  the  fall  and  winter  of 
1920-21. 

Study  of  the  concentration  of  sugar  in  the  blood  of  indi- 
vidual BIRDS.  It  is  a  common  practice  in  physiological  labora- 
tories to  keep  the  experimental  animals  for  only  a  short  time.  While 
there  are  probably  many  cases  where  repeated  observations  have 
been  made  on  individuals,  they  have  not  been  published  with  this 
point  in  view,  and  as  a  result  it  is  difficult  to  find  data  showing  whether 
or  not  the  concentration  of  the  sugar  in  the  blood  is  approximately 
constant,  or  whether  it  is  markedly  variable  from  time  to  time  in  one 
animal,  and  from  individual  to  individual. 

In  order  to  determine  these  points  for  the  pigeons,  two  were  set 
apart  to  be  used  exclusively  for  this  study.  They  were  kept  in  the 
laboratory  throughout  the  period  of  ol>servation  and  had  access  to 
food  and  water  at  all  times.     The  observations  were  extended  from 


BLOOD  SUGAR  OF  PIGEONS 


155 


October  29,  1920  until  the  death  of  pigeon  A  on  February  20,  and  until 
March  13,  1921  on  pigeon  B.  The  first  six  observations  were  made  at 
intervals  of  one  week.  Following  this  the  intervals  were  lengthened 
as  indictited  in  table  1. 

From  this  table  it  will  appear  that  pigeon  A,  in  all  but  four  of  the 
eleven  determinations,  had  a  sugar  content  of  175  mgm.  or  180  mgm. 
per  100  cc.  of  blood;  pigeon  B,  with  four  exceptions,  170  mgm.  or 
175  mgm.  per  100  cc.  These  variations  are  well  within  the  limits  of 
error  for  the  method,  and  so  are  to  be  considered  constant. 

TABLE  1 
Blood  sugar  of  normal  birds 


October  29... 
November  5. 
November  13 
November  20 
December  5 . . 
December  11. 
December  18, 
January  8 . . . . 
January  15 . . . 
February  6 . . . 
February  20 . 
March  13 


Weight 


grams 

317 
325 
327 
330 
330 
335 
335 
337 
340 
337 
342 


Glucose 
per  100  cc. 


mgm. 

200 
175 
175 
180 
150 
180 
175 
180 
175 
185 
195 


Weight 


grams 

368 
372 
375 
380 
377 
375 
370 
375 
377 
375 
377 
375 


Glucose 
per  100  cc. 


mgm . 

140 
170 
170 
175 
215 
175 
170 
170 
175 
165 
190 
175 


Obviously,  then,  there  is  a  concentration  of  sugar  which  is  charac- 
teristic of  the  blood  of  a  given  bird.  The  fact,  however,  that  the  two 
birds  which  happened  to  be  selected  yielded  characteristic  values  so 
close  together,  does  not  warrant  the  extension  of  this  value  to  other 
birds,  as  will  appear  later.  But  while  it  is  evident  that  there  is  a 
value  which  is  characteristic  of  a  given  bird,  it  is  also  evident  that 
this  value  will  not  necessarily  be  obtained  at  all  times.  The  occa- 
sional striking  variations  shown  in  the  table  illustrate  the  fact  that  here 
we  are  dealing  with  an  organism  which  is  responsive  to  modified  condi- 
tions, and  one  of  its  means  of  adjustment  to  its  environment  involves 
changes  in  the  concentration  of  blood  sugar.  This  offers  experimental 
confirmation  of  the  statement  of  Pike  and  Scott  (6)  that  the  concen- 
tration of  sugar  in  the  blood  is  one  of  those  internal  conditions  existing 


THE  AMERICAN  JOURNAL  OF  PHYSIOLOGY,  VOL.  58,  NO.  1 


156 


HANNAH    ELIZABETH   HONEYWELL 


in  higher  organisms  which  are  generally  constant,  and  which  are 
regulated  by  the  general  nervous  and  physico-chemical  mechanisms 
of  the  organism  as  a  whole. 

Three  cases  offered  an  opportunity  for  comparing  tlie  blood  sugar 
concentration  given  by  individuals  at  various  times  after  a  period  of 
48  hours'  inanition.  These  data  are  given  in  table  2,  from  which  it 
will  be  observed  that  each  bird  has  a  value  which  appears  to  be 
characteristic  for  it. 

TABLE  2 
Blood  sugar  of  individuals  after  inanition 


BLOOD  SUGAR  PER  100  CC. 

DATE 

Normal 

After  inanition 

rngvi. 

mgm. 

/ 

10-11 

190 

165 

'     1 

12-17 

175 

•  { 

10-22 

185 

165 

1-  7 

185 

20             1 

11-4 

140 

100 

12-18 

135      • 

TABLE  3 

Blood  sugar  of  normal  dogs  at  different  times 


DOG 

DATE 

BLOOD  SUGAR  PER  100  CC. 

mgm. 

^          { 

11-  6 

75 

1-  b 

73 

"          { 

10-31 

67 

11-  7 

65 

=          { 

11-20 

59 

1-16 

63 

These  results  are  similar  to  those  reported  by  Scott  and  Hastings 
(7)  for  dogs.  Two  determinations  each  were  made  on  three  different 
dogs,  the  greatest  difference  between  consecutive  determinations  upon 
the  same  dog  being  4  mgm.,  as  shown  in  table  3. 

In  a  series  of  22  blood  sugar  determinations  made  by  Kramer  and 
Cofl&n  (8)  on  a  quiet  dog,  the  amount  of  glucose  varies  from  87  mgm. 
to  93  mgm.  per  100  cc,  the  average  being  89  mgm.  per  100  cc. 


BLOOD    SUGAR    OF   PIGEONS 


157 


Jones  (9)  states  that  in  making  repeated  observations  on  individual 
rabbits  the  variations  were  within  the  experimental  error,  and  so  could 
be  considered  as  constant.  She  frequently  found,  however,  a  con- 
siderable variation  in  passing  from  individual  to  individual. 

In  a  recent  paper  Strouse  (10)  reported  a  series  of  observations  on 
five  normal  persons  covering  a  period  of  8  months.  In  the  series  the 
variations  for  individuals  range  from  27  mgm.  to  59  mgm.  per  100  cc. 
of  blood  with  an  average  variation  of  41  mgm.  per  100  cc.  Thallinger 
(11)  made  repeated  observations  on  a  boy  with  furunculosis,  whose 
blood  sugar  was  abnormally  high,  varying  from  145  mgm.  to  155  mgm. 
per  100  cc.  on  a  liberal  diet  which  was  low  in  carbohydrates. 

Thus  it  will  be  seen  that  the  pigeons  agree  with  the  rabbits  and  dogs 
in   possessing  a  characteristic  sugar  value.     Strouse 's  figures  would 

TABLE  4 
Effect  of  excitement  on    blood   sugar 


BIRD 

DATE 

BLOOD 

SUGAR  PER 

100  CC. 

REMARKS 

mgm. 

4 

/ 
I 

10-  9 
1-  7 

185 
265 

Excited  by  presence  of  several  strangers 

8 

{ 

10-22 
10-22 

185 
300 

Excited  by  loud  talking 

10 

{ 

11-19 
11-20 

175 
250 

Excited  by  escape  from  cage 

indicate  that  the  range  may  be  greater  in  man,  although  it  is  not  clear 
whether  this  greater  range  is  due  to  a  more  ready  response  to  changes 
in  the  environment  than  occurs  in  the  other  animals  studied,  or. 
whether  the  sugar-controlling  mechanism  is  not  so  perfectly  developed, 
or  whether  possibly  the  conditions  of  hving  were  not  so  carefully 
standardized 

Effect  of  handlixg;  emotional  glycemia.  As  noted  in  the 
foregoing,  noise,  loud  talking  and  the  presence  of  strangers  produce  a 
rise  in  the  blood  sugar.  Rough,  sudden  or  uncertain  handling  also 
disturbs  the  pigeon  and  increases  the  blood  sugar.  In  one  instance 
the  bird  escaped  from  the  cage  just  before  a  sample  was  drawn  and 
some  confusion  attended  its  recapture.  The  amount  of  sugar  in  the 
sample  of  blood  was  high,  as  shown  in  no.  10,  table  4. 


158  HANNAH    ELIZABETH   HONEYWELL 

From  this  table  it  will  be  seen  that  the  bird  offers  no  exception  to  the 
principle  long  ago  pointed  out  by  Boehm  and  Hoffman  (12),  Pavy 
(13)  and  others,  and  more  recently  by  Cannon  (14),  Shaffer  (15)  and 
Scott  (16),  that  to  obtain  blood  sugar  figures  of  value,  samples  of  blood 
must  be  obtained  without  pain  or  other  emotional  disturbance  of  the 
subject.  The  fact,  however,  that  birds  which  were  known  to  be  ex- 
cited have  such  high  figures  as  appear  in  table  4,  indicates  that  the 
lower  figure  of  about  185  mgm.  per  100  cc.  may  be  assumed  to  be  normal. 
This  was  the  value  obtained  by  Scott  and  Honeywell  (17)  and  though 
Fleming  (18)  found  a  much  lower  value  for  ducks,  in  fact  a  figure  quite 
comparable  with  that  characteristic  of  mammals,  the  normal  value  for 
the  pigeon,  at  least,  appears  to  be  much  higher  and  to  agree  well 
with  the  values  published  for  other  birds  (cf.  Scott  and  Honeywell). 

Effect  of  inanition.  For  reasons  which  will  be  discussed  later, 
the  birds  were  subjected  to  a  48-hour  period  of  inanition  in  determining 
the  alimentary  glycemia  curve  to  be  described  in  the  following  section. 
In  all  cases  the  concentration  of  sugar  in  the  blood  was  determined  soon 
after  the  arrival  of  the  birds  at  the  laboratory,  and  again  at  the  close 
of  the  48-hour  fast  and  just  before  feeding  the  glucose.  While  the 
effect  of  this  inanition  was  not  the  primary  purpose  of  the  experiment, 
this  procedure  offered  opportunity  for  its  study,  provided  that  the 
initial  values  can  be  taken  as  normal  values  for  birds  on  full  feed.  The 
propriety  of  this  is  in  some  doubt,  as  the  birds  had  not  yet  become  fully 
accustomed  to  their  new  environment  and  had  not  been  subjected  to 
standard  conditions.  This  would  probably  result  in  rather  wide 
variation  from  values  characteristic  for  the  individual  with  a  general 
tendency  to  yield  high  values.  It  is  felt,  however,  that  the  figures  as 
they   stand   merit   some   attention. 

As  noted  by  Rogers  (19),  the  general  effect  of  inanition  on  the  normal 
pigeon  is  to  increase  its  natural  restlessness.  It  becomes  irritable  and 
fights  on  the  slightest  provocation,  such  as  a  sudden  noise.  This 
might  lead  one  to  expect  higher  blood  sugar  values  in  birds  subjected 
to  inanition,  and  may  explain  those  values  which  are  even  higher  than 
normal  that  were  occasionally  found. 

From  table  5,  which  contains  the  data  for  the  blood  sugar  of  pigeons 
after  inanition,  it  appears  that  in  36  experiments  19  pigeons  show  a 
lower  blood  sugar  after  inanition  than  before.  The  results  vary  from 
3.1  per  cent  to  70  per  cent  below  the  initial  value.  Fifteen  pigeons  ex- 
hibited an  increase  in  blood  sugar  ranging  from  5  per  cent  to  100  per 
cent.     Two  pigeons  showed  no  change  whatever.     The  entire  series 


BLOOD    SUGAR    OF    PIGEONS 


159 


TABLE  5 

Effect  of  inanition 


WEIGHT 

BLOOD  SUGAR  PER  100  CC. 

DATE 

PERIOD 

OF 

INANITION 

BIRDS 

Before 
inani- 

After 
inani- 

Per 
cent  of 

Before 
inani- 

.^fter 
inani- 

Per cent 
of 

tion 

tion 

loss 

tion 

tion 

difference 

hours 

grams 

grams 

mgm. 

mgm. 

10 

November  17  to  19 

48 

280 

250 

10.7 

200 

175 

-  8.00 

11 

November  17  to  19 

48 

350 

320 

8.5 

175 

120 

-31.0 

12 

November  17  to  19 

48 

300 

240 

20.0 

155 

150 

-  3.1 

13 

December  15  to  17 

48 

300 

280 

6.6 

160 

100 

-37.6 

14 

December  15  to  17 

48 

310 

275 

11.3 

105 

90 

-14.2 

5 

December  15  to  17 

48 

340 

325 

4.4 

190 

175 

-  7.8 

9 

January  5  to  7 

48 

280 

272 

2.8 

140 

185 

32.0 

17 

January  5  to  7 

48 

325 

312 

4.0 

185 

265 

43.0 

18 

January  5  to  7 

48 

300 

290 

3.3 

175 

310 

77.0 

19 

January  5  to  7 

48 

350 

330 

5.6 

150 

290 

93.0 

20 

November  2  to  4 

48 

280 

248 

11.4 

140 

100 

-28.0 

21 

November  2  to  4 

48 

310 

280 

9.65 

120 

105 

-12.5 

22 

December  8  to  10 

48 

330 

305 

7.6 

200 

170 

-15.0 

23 

November  10  .to  12 

48 

340 

310 

8.8 

155 

155 

0.0 

14 

November  10  to  12 

48 

330 

315 

4.5 

105 

210 

100.0 

25 

November  10  to  12 

48 

230 

215 

6.5 

105 

170 

62.0 

26 

November  10  to  12 

48 

290 

250 

13.8 

255 

155 

-39.0 

27 

December  2  to  5 

48 

345 

340 

1.4 

120 

150 

25.0 

28 

December  2  to  5 

48 

330 

305 

7.6 

150 

140 

-  6.6 

29 

December  8  to  10 

48 

365 

340 

6.8 

120 

175 

46.0 

8 

December  16  to  18 

48 

3;3'0 

300 

9.1 

185 

55 

-70.0 

9 

December  16  to  18 

48 

300 

262 

12.6 

140 

135 

-  3.5 

4 

December  16  to  18 

48 

300 

290 

3.3 

185 

100 

-46.9 

33 

December  20  to  22 

48 

340 

330 

2.9 

160 

175 

9.4 

12 

December  20  to  22 

48 

330 

317 

3.9 

155 

155 

0.0 

11 

December  20  to  22 

48 

300 

285 

5.0 

175 

150 

-14.2 

36 

December  19  to  21 

48 

300 

280 

6.7 

185 

110 

-40.6 

37 

December  19  to  21 

48 

300 

285 

5.0 

190 

160 

-15.8 

38 

December  19  to  21 

48 

380 

340 

10.5 

185 

115 

-37.9 

9 

December  19  to  21 

48 

350 

340 

2.8 

140 

180 

28.6 

2 

September  11  to  14 

48 

220 

195 

11.3 

290 

300 

3.4 

3 

September  11  to  14 

48 

216 

200 

7.4 

190 

200 

5.2 

8 

October  20  to  22 

48 

356 

300 

15.7 

185 

200 

8.1 

9 

October  20  to  22 

48 

360 

280 

22.2 

187 

165 

-11.7 

51 

October  20  to  22 

48 

330 

272 

17.6 

195 

205 

5.1 

52 

November  2  to  4 

48 

340 

320 

5.9 

140 

160 

14.3 

Aver 

age 

166 

165 

-  0.6 

*^o^ • 

160  HANNAH    ELIZABETH   HONEYWELL 

gave  an  iaverage  decrease  in  blood  sugar  of  0.6  per  cent.  It  may, 
therefore,  be  that  48  hours  inanition  has  practically  no  effect  on  the 
blood  sugar  of  the  pigeon.  As  pointed  out  above,  the  evident  irri- 
tabihty  of  the  birds  subjected  to  inanition  with  its  possible  effect  upon 
the  concentration  of  sugar  in  the  blood  should  be  borne  in  mind. 

Effect  of  ingestion  of  glucose:  1.  Special  technique  and  dis- 
cussion. As  noted  in  the  previous  section,  the  sugar  was  determined 
upon  the  arrival  of  the  birds  in  the  laboratory.  Also  as  described 
above,  after  the  birds  had  become  accustomed  to  the  laboratory,  they 
were  subjected  to  a  fast  of  48  hours  and  the  sugar  in  the  blood  again 
determined.  The  results  of  these  two .  determinations  were  given  in 
table  5.  In  addition  to  the  reasons  usually  assigned  for  a  preliminary 
period  of  inanition,  this  somewhat  prolonged  period  seemed  to  be 
necessary  to  empty  the  crop  and  so  to  insure  a  rapid  passage  of  the 
sugar  to  the  region  of  the  alimentary  tract  where  absorption  might  be 
expected  to  take  place.  It  will  be  readily  appreciated  that  this  is  even 
more  essential  in  the  case  of  such  birds  as  the  pigeon,  which  are  pro- 
vided with  crop  and  gizzard,  neither  of  which  is  presumably  a  region 
of  absorption,  than  it  is  with  the  mammals,  and  possibly  than  it  would 
be  with  other  birds.  The  alimentary  canal  was  empty  in  all  birds 
examined,  so  this  period  of  inanition  may  be  considered  as  sufficient 
to  fulfill  its  purpose. 

After  the  second  sugar  determination,  the  appropriate  amount  of 
glucose  was  administered.  To  facilitate  the  feeding,  the  glucose  was 
made  into  tablets  and  a  weighed  amount,  1,  2  or  3  grams,  according  to 
the  series,  was  given  to  each  bird.  In  feeding  the  sugar,  the  beak  was 
opened  and  the  tablets  were  dropped  well  back  into  the  haouth.  If 
the  pellets  were  not  readily  swallowed,  a  little  water  was  given  through 
a  dropper.  Sometimes  gentle  stroking  of  the  throat  seemed  to  aid 
when   swallowing  wa§  especially   slow. 

Three  series  of  experiments  were  carried  out.  In  series  I,  each 
pigeon  was  fed  1  gram  of  glucose;. in  series  II,  2  grams;  and  in  series 
III,  3  grams.  In  terms  of  grams  per  kilogram  of  body  weight,  the 
average  amount  of  glucose  fed  was  4  grams,  7  grams,  and  10  grams 
in  the  respective  series. 

The  ordinary  clinical  test  for  carbohydrate  tolerance  is  100  grams  or 
about  1.4  grams  per  kilogram  of  body  weight,  if  the  average  weight  for 
man  is  taken  to  be  70  kilograms.  This  amount  was  fed  by  Cummings 
and  Piness  (20),  Hiller  and  Mosenthal  (21),  Jacobsen  (22),  Tachau  (23) 
and  Strouse,  who  has  also  fed  2  grams  and  2.8  grams  per  kilo  to  normal 
men.     Jones  gave  rabbits  an  average  dose  of  7.87  grams  per  kilogram. 


BLOOD  SUGAR  OF  PIGEONS 


161 


From  the  foregoing  it  will  be  seen  that  the  amounts  given  to  the 
pigeon  exceed  those  usually  given  man  in  similar  experiments.  In 
spite  of  this,  the  smallest  dose  used  in  the  present  experiments  which 
is  equivalent  to  one  of  1 .75  grams  for  a  man  weighing  70  kilograms,  had 
very  little  effect  on  the  concentration  of  sugar  in  the  blood  of  the 
pigeon.  In  a  man  such  an  amount  would  in  all  probability  raise  the 
concentration  of  blood  sugar  to  200  mgm.,  and  probably  induce  gly- 
cosuria. 

2.  Time  of  the  maximimi.  Since  it  was  desired  to  determine  the 
principal  points  in  the  entire  curve,  that  is,  to  follow  the  curve  to  its 
return  to  the  initial  value,  and  since  the  number  of  samples  of  blood 
which  could  be  drawn  safely  in  any  one  experiment  was  limited  be- 
cause of  the  injury  to  the  heart  which  would  result  from  repeated 

TABLE  6 
Time  of  maximum  of  alimentary  glycemia 


BLOOD  SUGAR 

BLOOD  SUGAR  PER  100  CC.  AFTER  FEEDING  GLUCOSE 

BIRD 

GLUCOSE  FED 

AFTER 
INANITION 

1  hour 

2  hours 

3  hours 

4  hours 

5  hours 

grams 

mgm. 

mgm. 

mgm. 

mgm. 

mgm. 

61 

3 

160 

155 

240 

400 

335 

320 

62 

3 

180 

185 

235 

305 

210 

215 

64 

2 

210 

205 

245 

315 

275 

260 

65 

2 

185 

200 

215 

240 

250 

205 

66 

1 

170 

190 

200 

215 

240 

200 

67 

1 

165 

170 

185 

205 

225 

190 

punctures  at  short  intervals,  it  was  necessary  to  determine  the  time 
elapsing  between  the  administration  of  the  glucose  and  the  maximum 
sugar  concentration  in  the  blood.  For  this  purpose,  as  indicated  in 
table  6,  hourly  determinations  were  made  after  the  sugar  was  fed.  It 
will  be  seen  from  the  results  given  in  this  table  that  the  maximum  may 
be  assumed  to  occur  between  the  third  and  -fourth  hours.  This  last 
interval  was  therefore  allowed  to  elapse  after  feeding  and  before  drawing 
the  first  sample,  and  the  sugar  level  at  this  time  may  be  assumed  very 
nearly  to  represent  the  maximum  attained. 

When  3  grams  of  glucose  were  fed  the  maximum  occurred  at  or  about 
the  third  hour.  When  2  grams  of  glucose  were  fed  the  maximum 
occurred  in  one  case  at  the  third  hour  and  in  the  other  case  at  the  fourth 
hour.  After  the  feeding  of  1  gram  of  glucose,  the  maximum  occurred 
at  about  the  fourth  hour.     From  these  results,  given  in  table  6,  it  will 


162 


HANNAH    ELIZABETH   HONEYWELL 


appear  that  the  greater  the  amount  of  glucose  fed  the  earlier  the  maxi- 
mum will  be  reached. 

In  this  connection  it  is  interesting  to  note,  as  Strouse  has  pointed  out, 
that  a  heavy  dosage  often  has  the  effect  in  man  of  delaying  the  onset  of 
the  maximum  rather  than  accelerating  it  as  in  the  pigeon.  There 
must  be,  then,  some  fundamental  difference  between  the  carbohydrate 
economy  of  the  pigeon  and  that  of  man. 

3.  Course  of  alimentary  hyperglycemia.  In  each  case  samples  of 
blood  were  drawn  just  before  the  administration  of  the  glucose  and 
again  after  the  lapse  of  4,  6  and  24  hours.     The  results  are  collected  in 


J.GO 


l¥ 


tables  7,  8  and  9  and  summarized  in  the  accompanying  curves  (fig.  1). 
In  the  first  series  one  gram  of  glucose  was  given  each  bird.  As  noted 
before,  this  is  about  double  the  ratio  of  the  test  meal  usually  given 
man  for  diagnostic  purposes.  From  table  7  it  will  appear  that  there  is 
no  change  or  as  occurs  more  frequently,  only  a  slight  rise  at  the  end  of 
the  first  period.  The  average  for  the  series  gives  a  rise  of  18  per  cent 
at  this  time. 

In  the  second  and  third  series  there  is  a  different  manifestation.  In 
the  second  series  2  grams,  and  in  the  third  3  grams  were  given  each  bird. 

The  average  rise  at  the  end  of  4  hours  in  the  second  series  was  45  per 
cent,  and  in  the  third,  93  per  cent.     From  table  8  it  will  be  seen  that 


BLOOD    SUGAR    OF    PIGEONS 


163 


m  the  second  series  only  five  birds  had  returned  to  their  previous  level 
at  the  end  of  24  hours;  and  in  the  third  series,  table  9,  one  alone  had 
returned  in  that  interval  to  the  level  which  existed  before  the  ingestion 
of  the  glucose. 

While  the  average  at  the  end  of  the  inanition  period  may  vary  some- 
what for  the  different  groups,  the  average  at  the  end  of  24  hours  after 
feeding  the  glucose  is  approximately  that  of  the  normal  birds.  Be- 
cause of  the  relatively  low  initial  values  found  in  the  second  and  the 
third  series,  the  final  values  found  for  these  series  are  distinctly  higher 

TABLE  7 
Effect  of  ingestion  of  glucose  upon  the  sugar  in  the  blood 

Series  I 


GLU- 

BLOOD SUGAR  PER  100  CC. 

COSE 

AFT  EI 

I     FEEDING  GLUCOSE 

BIRD 

DATE 

WEIGHT 

FED 
PER 
KILO 

OF  IN- 
CREASE 

REMARKS 

Inani- 
tion 

4 
hours 

6 
hours 

24 
hours 

grams 

(jrams 

mgm. 

mgm. 

mgm. 

mgm. 

10 

November  19 

250 

4.0 

175 

100 

110 

250 

-37.0 

Bird  excited  by 

11 

November  19 

320 

3.1 

120 

150 

150 

125 

25.0 

escape  from 

12 

November  19 

240 

4.2 

155 

150 

160 

150 

3.2 

cage  before 

13 

December  17 

■  280 

3.5 

100 

275 

245 

195 

175.0 

drawing  of  24 

14 

December  17 

275 

3.6 

90 

210 

300 

270 

233.0 

hour  sample 

5 

December  17 

325 

3.1 

175 

170 

160 

230 

-  2.8 

9 

January  7 

272 

3.7 

185 

270 

175 

265 

46.0 

17 

January  7 

312 

3.2 

265 

280 

275 

275 

5.7 

18 

January  7 

290 

3.4 

310 

350 

300 

250 

1.3 

19 

January  7 

330 

3.0 

290 

295 

260 

155 

1.7 

Average 

259 

3.9 

190 

225 

175 

180 

18.0 

than  the  initial  values.  Whether  or  not  this  is  significant  we  are  not 
prepared  to  state  definitely,  although  it  would  seem  to  be  accidental. 
Unfortunately  there  is  very  little  data  available  which  permits  of  a 
comparison  of  the  course  of  the  alimentary  hyperglycemia  of  different 
species.  In  fact,  only  three  species  seem  to  have  been  studied  from 
this  point  of  view.  In  her  recent  paper,  Jones  has  made  determinations 
on  the  blood  of  rabbits  only  at  a  single  period  after  the  ingestion  of  the 
glucose.  From  the  work  of  Bang  (24),  the  1-hour  period  which  she 
chose  would  presumably  give  figures  at  or  near  the  maximum  attained. 
Her  results  do  not,  however,  permit  one  to  follow  the  course  of  the  curve. 
Bang  reported  a  short  series  of  experiments  on  rabbits  which  had  been 


164 


HANNAH    ELIZABETH    HONEYWELL 


fed  from  5  to  20  grams  of  glucose.  After  a  S-daj^  period  of  inanition  he 
found  that  the  maximum  was  reached  in  1|  to  2^  hours,  and  that  in 
general  the  sugar  level  had  returned  to  normal  in  6  hours.  The  amount 
of  sugar  did  not  seem  materially  to  modif}^  the  time  relations  of  the 
curve.  The  same  may  be  said  of  a  similar  but  even  shorter  series, 
in  which  sugar  was  given  without  a  previous  period  of  inanition. 

Fisher  and  Wishart  (25)  in  experimenting  with  dogs  weighing  8  to 
9  kilograms,  fed  approximately  6  grams  of  glucose  per  kilogram,  and 
found  that  the  maximum  blood  sugar  occurred  one  hour  after  the 
ingestion  of  the  glucose. 

TABLE  8 
Series  II 


BIRD 

DATE 

WEIGHT 

GLUCOSE 

FED  PER 

KILO 

BLOOD  SUGAR  PER  100  CC.  . 
FEEDING  GLUCOSE 

A.FTER 

PER  CENT 
OF  IN- 

Inani- 
tion 

4  hours 

6  hours 

24  hours 

CREASE 

grams 

grams 

Tngm. 

mgm. 

mgm. 

mgm. 

20 

November  4 

248 

8.4 

100 

150 

125 

295 

50.0 

21 

November  4 

280 

7.1 

105 

300 

250 

180 

185.0 

22 

December  10 

305 

6.5 

170 

290 

280 

2b0 

70.0 

23 

November  12 

310 

6.4 

155 

440 

140 

145 

184.0 

14 

November  12 

315 

6.3 

210 

210 

■    250 

240 

19.0 

25 

November  12 

215 

9.3 

170 

200 

190 

120 

17.0 

26 

November  12 

250 

8.0 

155 

250 

210 

160 

61.0 

27 

December  4 

340 

5.9 

150 

280 

100 

130 

87.0 

28 

December  4    . 

305 

6.5 

140 

200 

285 

175 

103.0 

29 

December  10 

340 

5.9 

175 

270 

160 

110 

54.0 

Aven 

lee 

290 

6.9 

165 

240 

195 

180 

45.0 

The  work  of  many  investigators,  notably  Cummings  and  Piness, 
Hiller  and  Mosenthal,  Hamman  and  Hirschman  (26),  Jacobsen  and 
Strouse,  indicates  that  after  the  ingestion  of  100  grams  of  glucose  the 
maximum  concentration  of  sugar  in  the  blood  of  man  occurs  normally 
in  about  30  minutes  and  that  it  has  returned  approximately  to  its 
previous  level  by  the  end  of  the  second  hour.  Jacobsen  and  Strouse 
point  out  that  occasionally  the  maximum  is  attained  only  after  a  longer 
period;  and  that  when  this  is  true  the  level  is  apt  to  be  higher  than  usual, 
and  the  return  to  the  previous  value  is  usually  slower. 

Strouse  particularly  calls  attention  to  the  fact  that  in  diabetes  and 
other  conditions  which  may  be  presumed  to  alter  the  carbohydrate 
metabolism,  such  curves  are  common  but  that  such  individuals  may  be 


BLOOD   SUGAR    OF   PIGEONS 


165 


induced  to  give  the  ''normal"  or  usual  curve  if  given  less  sugar.  On 
the  other  hand  normal  individuals  will  give  the  ''diabetic"  curve  if 
the  dose  be  doubled  or  tripled. 

A  study  of  the  curves  obtained  from  pigeons  shows  that  the  maximum 
occurs  from  the  third  to  the  sixth  hour  after  feeding,  and  when  amounts 
were  fed  which  essentially  altered  the  sugar  level,  the  curve  did  not 
return  to  normal  for  a  much  longer  period,  in  some  cases  exceeding 
2-4  hours.  Thus  they  resemble  more  closely  the  delayed  curves  ob- 
tained from  men  rather  than  the  usual  or  normal  one,  and,  at  first 
thought,  the  obvious  reason  is  the  very  heavy  dose  of  glucose  given  to 
the  birds. 

TABLE  9 
Series  III 


GLU- 
COSE 

BLOOD  SuUaR  per  100  CC. 
AFTER  FEEDING  GLUCOSE 

PER 

1 

BIRD 

DATE 

WEIGHT 

FED 
PER 
KILO 

1 

OF  IN- 
CREASE 

REMARKS 

Inani- 
tion 

4 
hours 

6 
hours 

mgm. 

24 
hours 

grams 

grams 

mgm. 

mgm. 

mgm. 

8 

December  18 

300 

10.0 

55 

215 

200 

205 

291.0 

20 

December  18 

262 

11.4 

135 

320 

150 

160 

137.0 

Excited  by    pres- 

4 

December  18 

290 

10.3 

100 

160 

300 

22Q 

200.0 

ence  of  strangers 

33 

December  20 

330 

9.0 

175 

335 

200 

110 

91.5 

when  third  sam- 

12 

December  20 

317 

9.4 

155 

240 

210 

225 

55.0 

ple  was  shown. 

11 

December  20 

285 

10.5 

150 

320 

195 

175 

113.0 

Struggled  and  died 

36 

December  21 

280 

10.7 

110 

225 

205 

260 

104.5 

during  drawing 

37 

December  21 

285 

10.5 

160 

200 

265 

180 

65.5 

of  last  sample. 

38 

December  21 

340 

8.8 

115 

265 

110 

195 

130.4 

9 

December  21 

340 

8.8 

180 

330 

220 
205 

210 

180 

84.0 

Average 

303 

9.8 

1 

135 

260 

93.0 

In  order  to  determine  this  point,  that  is,  whether  the  pigeons  would 
respond  to  a  smaller  dose  and  whether  or  not  a  maximum  occurring 
during  the  first  hour  had  been  overlooked,  four  birds  were  fed  0.4 
gram  of  glucose  each,  after  an  inanition  period  of  48  hours.  This  is 
the  amount  of  glucose  which,  for  the  weight  of  the  bird,  approximates 
the  usual  test  dose  for  man.  Blood  sugar  determinations  were  made 
30  minutes,  1  hour  and  If  hours  after  the  ingestion  of  the  glucose.  The 
results  are  shown  in  table  10.  Since  the  variations  were  all  within  the 
limits  of  experimental  error,  it  may  be  concluded  that  the  pigeon  does 
not  respond  to  as  small  a  dose  as  does  man  and  that  there  is  a  fundamen- 
tal reason  for  the  difference  in  the  alimentary  glycemia  curves  shown 
by  the  two  species. 


166 


HANXAH    ELIZABETH   HONEYWELL 


In  his  series  on  normal  men  Strouse  obtained  an  average  increase  of 
42  per  cent  after  an  ingestion  of  100  grams  of  glucose,  or  1.4  grams  per 
kilogram.  The  pigeons  show  an  average  increase  of  only  18  per  cent 
after  the  ingestion  of  1  gram  or  about  3.5  grams  per  kilogram,  and  it 
was  not  until  they  had  been  given  2  grams  or  7  grams  per  kilogram  that 
they  approached  the  percentage  increase  reported  by  Strouse  for  men. 

In  addition  it  should,  perhaps,  be  pointed  out  that  the  resemblance 
is  more  apparent  than  real  for,  as  mentioned  above,  the  effect  of  the 
size  of  the  dose  upon  the  time  elapsing  between  the  administration  of 
the  dose  and  the  occurrence  of  the  maximum  is  in  the  opposite  sense 
in  the  pigeon  and  in  man.  It  would  thus  seem  that  the  mechanism 
of  storage  of  sugar  is  somewhat  different  in  the  two  groups. 

Post-mortem  examinations  of  two  pigeons  which  were  killed  after 
inanition  and  before  feeding  showed  that  the  crop  and  gizzard  were 

TABLE  10 

Effect  of  varying  amounts  of  glucose  on  time  of  maximum 


BIRDS 

WEIGHT 

AMOUNT  OF 
GLUCOSE  FED 

BLOOD  SUGAR  IN  MGM.  PER  100  CC. 
FEEDING  GLUCOSE 

AFTER 

PER  KILO 

Inanition 

h  hour 

1  hour 

U  hours 

grams 

grams 

1 

250 

1.6 

180 

165 

180 

175 

2 

370 

1.1 

220 

225 

230 

225 

3 

280 

1.4 

195 

200 

190 

205 

4 

350 

1.1 

185 

175 

180 

175 

empty,  while  the  intestine  contained  only  a  small  amount  of  fluid. 
Conditions  were  the  same  in  pigeons  which  were  examined  at  the  end 
of  the  fourth  and  sixth  hour  after  feeding.  The  contents  of  the  ali- 
mentary tract  were  not  tested  for  the  presence  of  sugar.  Conse- 
quently, while  the  indications  as  they  stand  are  that  the  delay  in 
reaching  the  maximum  is  not  due  to  delay  in  absorption,  but  rather  to 
some  peculiarity  in  the  mechanism  of  storage,  one  is  not  justified  in 
definitely  drawing  such  a  conclusion  until  a  study  of  the  contents  of  the 
alimentary  canal  has  been  made  in  parallel  with  blood  sugar  deter- 
minations. 

Since  concentration  of  the  sugar  in  the  bird  is  normally  so  high  as 
compared  with  mammals,  in  this  particular  resembling  the  diabetic, 
and  since  the  curve  obtained  from  birds  somewhat  resembles  that  ob- 
tained from  diabetic  man,  there  may  possibly  be  some  relationship  be- 


BLOOD    SUGAR    OF    PIGEONS  167 

tween  the  absolute  initial  height  of  the  sugar  concentration  and  the 
form  of  the  curve  of  alimentary  hyperglj^cemia.  However,  as  noted 
above,  it  would  seem  more  probable  that  in  the  birds  the  storage 
mechanism  is  somewhat  different  from  that  common  in  mammals 
and  that  further  work  must  be  done  before  a  satisfactory  correlation  is 
possible. 

SUMMARY 

1.  Each  bird  has  a  characteristic  sugar  level  about  which  it  varies 
from  day  to  day.     In  this  it  resembles  the  rabbit  and  dog. 

2.  These  individual  variations  are  caused  by  variations  in  the  ex- 
ternal and  internal  environment  of  the  bird. 

3.  A  series  of  inanition  values  for  blood  sugar  is  given  and  com- 
pared with  the  values  found  on  full  diet.  From  these  figures  it  is 
concluded  that  48  hours'  inanition  has  practically  no  effect  on  the  blood 
sugar  of  the  pigeon. 

4.  It  has  been  found  that,  in  general,  when  from  1  to  3  grams  of 
glucose  are  fed  to  the  pigeon  the  maximum  rise  in  the  blood  sugar 
occurs  in  3  to  4  hours. 

5.  It  is  indicated  that  the  greater  the  amount  of  glucose  given  the 
earlier  will  the  maximum  be  reached. 

6.  When  1  gram  of  glucose  or  less  is  fed  to  pigeons  there  is  very 
little  modification  of  the  sugar  in  the  blood.  When  2  or  3  grams  are 
fed  there  is  a  manifest  rise  in  the  blood  sugar  which  gradually 
approaches  its  former  level. 

The  author  wishes  to  acknowledge  her  indebtedness  to  Prof.  E.  L. 
Scott  for  his  assistance  in  the  execution  of  this  work. 


BIBLIOGRAPHY 

Warburg:  Hoppe-Seyler's  Zeitschr.,  1910,  Ixvi,  305;  Ixxvi,  331. 

Bierry:  Comptes  Rendus,  1919,  clxix,  1112. 

Hollinger:  Deutsche.  Arch.  f.  klin.  Med.,  1908,  xcii,  217. 

Mosso:  Fatigu,  New  York,  1904. 

MacLean:  Biochem.  Journ.,  1919,  xiii,  135. 

Pike  and  Scott:  Amer.  Naturalist,  1915,  xHv,  321. 

Scott  and  Hastings:  Proc.  Soc.  Exper.  Biol,  and  Med.,  1920,  xvii,  67. 

Kramer  and  Coffin:  Journ.  Biol.  Chem.,  1916,  xxv,  423. 

Jones:  Journ.  Biol.  Chem.,  1920,  xliii,  507. 

Strouse:  Arch.  Int.  Med.,  1920,  xxvi,  751. 

Thallinger:  Journ.  Amer.  Med.  Assoc,  1921,  Ixxvi,  295. 


168  HANNAH    ELIZABETH   HONEYWELL 

(12)  BoEHM  AND  Hoffmann:  Arch,  f,  Exper.  Path.  u.  Pharm.,  1878,  viii,  271. 

(13)  Pavy:  Journ.  Physiol.,  1899,  xxiv,  479. 

(14)  Cannon  :  This  Journal,  1914,  xxxiii,  356. 

(15)  Shaffer:  Journ.  Biol.  Chem.,  1914,  xix,  285. 

(16)  Scott:  This  Journal,  1914,  xxxiv,  271. 

(17)  Scott  and  Honeywell:  This  Journal,  1921,  Iv,  362. 

(18)  Flemming:  Journ.  Physiol,  1920,  liii,  236. 

(19)  Rogers:  This  Journal,  1916,  xli,  555. 

(20)  CmiMiNGS  AND  PiNESs:  Arch.  Int.  Med.,  1917,  xix,  777. 

(21)  HiLLER  AND  MosENTHAL :  Joum.  Biol.  Chem.,  1917,  xxviii,  197. 

(22)  Jacobsen:  Biochem.   Zeitschr.,   1913,  Ivi,  471. 

(23)  Tachau:  Arch.  f.  klin.  Med.,  1911,  civ,  437. 

(24)  Bang:  Der  Blutzucker,  1913. 

(25)  Fisher  and  Wishart:  Journ.  Biol.  Chem.,  1912,  xiii,  49. 

(26)  Hamman  and  Hirschmann:  Arch.  Int.  Med.,  1917,  xx,  761. 


VITA 

Hannah  Elizabeth  Honeywell  was  born  in  Walton,  N.  Y.,  April  3, 
1888.  She  graduated  from  Walton  High  School  in  1906,  received 
the  degree  of  Batchelor  of  Arts  from  Mount  Holyoke  College  in  1910, 
and  the  degree  of  Master  of  Arts  from  Columbia  in  1917. 

Her  publications  are: 

A  Study  of  the  Sugar  in  the  blood  of  Normal  Pigeons.  (With 
E.  L.  Scott.)     American  Journal  of  Physiology,  1921,  Iv,  362. 


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